Graded or discrete? A quantitative analysis of Campbellʹs monkey alarm calls

A standard way of describing the vocal behaviour of nonhuman primates is to classify the vocal repertoire as either graded or discrete. We analysed a large database of calls given by adult males of a primate considered a typical example for discrete vocal behaviour, the forest-dwelling Campbellʹs monkeys, Cercopithecus campbelli. We recorded vocal responses from several dozen individuals to their main predators, crowned eagles and leopards. Using cluster analysis techniques, we found two main call types, which were modified further by optional affixation of an inflexible vocal structure. It was possible to force the four call types into eight subtypes, with various degrees of gradedness. When taking context into account, we found that acoustically discrete and nonaffixed calls tended to be given right after discovering a predator, while acoustically graded and affixed calls were given during later parts of a predator encounter and to nonpredatory disturbances. In sum, our results suggest that classifications of primate vocal repertoires as either discrete or graded are likely to be meaningless, as communicatively relevant acoustic variation can be present within seemingly discrete call types.